When I visited Dinosaur National Monument in October with Brian Engh and Yara Haridy, we spent a decent amount of time checking out DNM 28, a skull and associated bits of Camarasaurus. In particular, I got some shots of the axis (the second cervical vertebra behind the head), and it got me thinking about pneumaticity in this unusual element. Why I failed to get a full set of orthogonal shots is quite beyond my capacity, but we can roll with what I have. Before we go on, you might want to revisit Tutorial 36 to brush up on the general parts of the atlas-axis complex.

Here’s the axis in left lateral view (so, anterior to the left).

And a labeled version of the same. A few things to note:

• One oddity of sauropod axes (and of axes of most critters) is that not only are the articular facets of the prezygapophyses not set forward of the neural arch, they’re set backward, well behind the forward point of the arch.
• The dens epistrophei or odontoid process is sticking out immediately below the neural canal. This is the tongue of bone that articulated with the atlas (first cervical vertebra) in life.
• Check out the prominent epipophysis above the postzygapophysis, which anchored the long dorsal neck muscles. (For more on epipophyses, see these posts, and especially this one.)
• The diapophysis and parapophysis articulated with a cervical rib, which is not shown here. In fact, I don’t remember seeing it in the drawer that this vert came from. The atlantal and axial cervical ribs are small, apparently fused late in life if they fused at all, and are easily lost through taphonomic processes.
• At least three pneumatic features are visible in this lateral view: the lateral fossa on the centrum, which is referred to as the “pleurocoel” in a lot of older literature; a ventral fossa that lies between the parapophysis and the midline ventral keel; and a fossa on the neural arch, behind the postzygodiapophyseal lamina. In the nomenclature of Wilson et al. (2011), this is the postzygocentrodiapophyseal fossa.

“Postzygocentrodiapophyseal fossa” is a mouthful, but I think it’s the only way to go. To be unambiguous, anatomical terminology needs to references specific landmarks, and the schemes proposed by Wilson (1999) for vertebral laminae and Wilson et al. (2011) for vertebral fossae are the bee’s knees in my book.

Nomenclatural issues aside, how do we know that these fossae were all pneumatic? Well, they’re invasive, there’s no other soft-tissue system that makes invasive fossae like that in archosaur vertebrae (although crocs sometimes have shallow fossae that are filled with cartilage or fat), and the same fossae sometimes have unambiguous pneumatic foramina in other vertebrae or in other sauropods.

Most of the features labeled above are also visible on the right side of the vertebra, although the ventral fossa is a little less well-defined in this view, and I can’t make out the prezyg facet. Admittedly, some of the uncertainty here is because of my dumb shadow falling across the vertebra. Specimen photography fail!

The paired ventral fossae are more prominent in this ventral view, on either side of the midline ventral keel (anterior is to the top).

And here’s a labeled version of the same ventral view.

Finally, here’s the posterior view. It’s apparent now that the neural spine is a proportionally huge slab of bone, like a broad, tilted shield between the postzygapophyses (which are also quite large for the size of this vertebra). The back side of the neural spine is deeply excavated by a complex fossa with several subfossae (kudos again to Jeff Wilson [1999] for that eminently useful term).

Here’s the same shot with some features of interest labeled. If I’ve read Wilson et al. (2011) correctly, the whole space on the back side of the neural spine and above the postzygs could be considered the spinopostzygapophyseal fossa, but here I’ve left the interspinous ligament scar (ILS) unshaded, on the expectation that the pneumatic diverticula that created that fossa were separated on the midline by the interspinous ligament. I might have drawn the ILS too conservatively, conceivably the whole space between the large deeply-shadowed subfossae was occupied by the interspinous ligament.

I’m particularly interested in those three paired subfossae, which for convenience I’m simply calling A, B, and C. Subfossa A may just be the leftover space between the spinopostzgyapophyseal laminae laterally and the interspinous ligament medially. I think subfossa B is invading the ramus of bone that goes to the epipophysis and postzygapophysis, but I didn’t think to check and see how far it goes (that might require CT anyway).

Subfossa C is the most intriguing. Together, those paired fossae form a couple of shallow pits, just on either side of the midline, and aimed straight forward. They can’t be centropostzygapophyseal fossae, which used to be called peduncular fossae, because they’re not in the peduncles on either side of the neural canal, they’re up above the lamina that connects the two postzygapophyses. Could they be ligament attachments? Maybe, but I’m skeptical for at least four reasons:

1. Although interspinous ligament attachments often manifest as pits in the cervical vertebrae of birds, in sauropods they usually form rugosities or even spikes of bone that stick out, not inward. Furthermore, these pits are smooth, not rough like the interspinous ligament scars of birds.
2. The interspinous ligament in tetrapods is typically a single, midline structure, and these pits are paired.
3. Similar pits in front of the neural spine are present in some sauropod caudals, and they appear to be pneumatic (see Wedel 2009: p. 11 and figure 9).
4. Pits at the base of the neural spine seem to be fairly uncommon in sauropod vertebrae. If they were attachment scars from the universally-present interspinous ligaments, we should expect them to be more prominent and more widespread.

But if these paired pits are not ligament scars, what are they? Why are they present, and why are they so distinct? Sometimes (often?) subfossae and accessory laminae look like the outcome of pneumatic diverticulum and bone reacting to each other (I almost wrote ‘playing together’), in what looks like a haphazard process of adaptation to local loading. But the symmetry of these pits argues against them being incidental or random. They don’t seem to be going anywhere, so maaaybe they are the first hoofbeats of the embossed laminae and “unfossae” that we see in the vertebrae of more derived sauropods (for which see this post), but again, their symmetry in size and placement isn’t really consistent with the “developmental program gone wild” appearance of “unfossae”. I really don’t know what to make of them, but if you have ideas, arguments, or observations to bring to bear, the comment field is open.

In summary, sauropod axes are more interesting than I thought, even in a derpasaurus like Cam. I’ll have to pay more attention to them going forward.

References

• Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.
• Wilson, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 19(4): 639-653. DOI: 10.1080/02724634.1999.10011178
• Wilson, J. A., Michael, D. D., T. Ikejiri, E. M. Moacdieh, and J. A. Whitlock. 2011. A nomenclature for vertebral fossae in sauropods and other saurischian dinosaurs. PLoS One 6(2): e17114. DOI: 10.1371/journal.pone.0017114